LESSON 5

Tissues and Organs; or how the Plant is built

(by Alexey Shipunov)

5.1 Tissues

From now on, we will frequently use multiple names of plants2 group, they are
summarized on Figure 5.1, and in more details — on Fig. 6.1.

Figure 5.1. Plants2 classification: overview.

5.1.1 Epidermis and Parenchyma

Why did plants go on land? In order to escape competition with other plants for
resources like the sun and nutrients, but also to obtain much more sunlight that
was otherwise seriously reduced underwater. The move to land also helped plants
escape predators. Lastly, plants benefited from this change because they
escaped from the temperature-gases conflict: warmer temperatures are good
for organisms but significantly decrease the amount of gases diluted in water.

Although this action solved several problems, it also raised new issues that
needed to be dealt with. The most important was the risk of drying out. To
combat this, plants developed their first tissue: epidermis covered with a cuticle
which served a purpose similar to a plastic bag. For the really small (millimeters)
plant it is enough because, in accordance to surface / volume law, they have high
relative surface, and diffusion can serve for gas exchange.

However, bigger plants also need to exchange gases, and they developed stomata
which served as a regulated pore system. The remaining cells became second
tissue: parenchyma (or ground tissue, or main tissue).

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Another response (Fig. 5.3) for drying was a development of poikilohydricity (see
below), the ability to hibernate in (almost) dried condition. As hibernation is
generally dangerous since it requires “system restart”, that evolutionary route
did not become the main.

Tissue is a union of cells which have common origin, function and similar
morphology. Tissues belong to organs: organ is a union of different tissues which
have common function(s) and origin. Plants have simple and complex tissues. The
simple tissues are composed of the same type of cells; complex tissues are
composed of more than one type of cell, these are unique to plants.

Parenchyma (Fig. 5.4) are spherical, elongated cells with a thin primary cell wall.
It is a main component of young plant organs. The basic functions of parenchyma
are photosynthesis and storage. Parenchyma cells are widespread in plant body.
They fill the leaf, frequent in stem cortex and pith and is a component of complex
vascular tissues. Contrary to parenchyma (which is a simple tissue), epidermis is a
complex tissue composed of epidermal and stomata cells. Its main functions are
transpiration, gas exchange and defense.

As it seen here, plants acquired tissues in a way radically different from animals
(Fig.5.2) : while plants regulate gas and water exchange in response to terrestrial
environment, animals actively hunt for food (using kinoblast tissues) and then
digest it (with pagocytoblast tissue).

5.1.2 Supportive Tissues: Building Skyscrapers

When more and more plants began to move from the water to the land,
competition once again became a problem (Fig. 5.3). To solve this, plants followed
“Manhattan solution”: they grew vertically in order to be able to escape
competition for the sunlight and therefore must develop supportive tissues.

Figure 5.2. Phagocytella (proto-animal) with kinoblast and phagocytoblast vs. proto-plant with
epidermis and ground tissue.

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Collenchyma (Fig. 5.4) is living supportive tissue that has elongated cells and a
thick primary cell wall. Its main function is the mechanical support of young stems
and leaves via turgor.

Sclerenchyma (Fig. 5.4) is a dead supportive tissue that consists of long fibers or
short, crystal-like cells. Each cell has a thick secondary wall that is rich in lignin.
Its main function is a support of older plant organs, and also hardening different
parts of plants (for example, make fruit inedible before ripeness so no one will
take the fruit before seeds are ready to be distributed). Without sclerenchyma,
if a plant isn’t watered, the leaves will droop because the vacuoles will decrease in
size which lowers the turgor. Fibers inside phloem (see below) are sometimes
regarded as a separate sclerenchyma.

Three times in their evolution plants found the new application for lignin or
similar polymers: at first, similar chemicals covered the spore wall which was an
adaptation to the spore distribution with wind. Then similar chemicals were used
to make cuticle, “epidermal plastic bag” to prevent transpiration outside of
stomata. Finally, with acquiring of sclerenchyma, plants found how to use dead
cells with completely lignified cell walls.

By the way, stomata likely had a similar fate, they historically appeared on
sporangia to help them dry faster and release spores effectively. Regulation of
transpiration is their second function.

Figure 5.3. Challenges

to land plants and their
responses, part 1. Part 2
is on Fig. 7.14.

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Figure 5.4. Left to right, top to bottom: parenchyma, sclerenchyma (cross- and longitudinal
sections) and collenchyma. First three photos from the stem of Helianthus, fourth from Medicago
stem. Magnification ×400.

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Cell types and tissues

“Parenchyma” and “sclerenchyma” terms are freShoot systemquently used in two

ways: first, to name tissues (or even classes of tissues) which occur in multiple
places of the plant body, and second, to name the cell types which are components
of tissues. Therefore, it is possible to say “parenchyma of stem”, “parenchyma of
stem pith”, “parenchyma of xylem” and even “leaf mesophyll is a parenchyma”.

5.1.3 Meristems: the Construction Sites

Plant growth requires centers of development which are meristems. Apical

meristems are centers of plant development located on the very ends of roots
(RAM) and stems (SAM). They produce intermediate meristems (like procambium)
which form all primary tissues. The lateral meristem or cambium originates from
the procambium which in turn originates from apical meristems. It usually arises
between two vascular tissues and its main functions are thickening and producing
secondary vascular tissues (Fig. 5.5).

Figure 5.5. Meristems and tissues.

Other meristems include: intercalary which elongate stems from the “middle”,
marginal which are responsible for leaf development and repair meristems arising
around wounds, they also control vegetative reproduction.

5.1.4 Vascular Tissues

Bigger plants escaped from competition and performed effective metabolism.

However, with all the growth the plants went through, their size became too big
for slow symplastic plasmodesmata connections. Another, filter paper-like
apoplastic transport was also not powerful enough. The solution was to develop
vascular tissues, xylem and phloem (Fig. 5.6, Fig. 5.29).

The main functions of xylem are the transportation of water and mechanical
support. The xylem may be found either in a vascular bundle or a vascular cylinder.
The three types of xylem cells are tracheary elements (these include tracheids
and vessel members), fibers, and parenchyma. Xylem elements, except for the
parenchyma, are rich in lignin and are main components of wood. Tracheids are
closed on both ends and connected with pits whereas vessel members are more or

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less open and connects via perforations. Tracheids, vessel members and fibers ]
are dead cells. Xylem parenchyma, on the other hand, is alive.

Figure 5.6. Cells of xylem (left, a–d) and phloem (right, e–h): a fibers, b vessels with open
perforations, c parenchyma, d tracheids with pits, e parenchyma, f fibers, g sieve tubes, h
companion cells.

Pits of tracheids consist of a pit membrane and the torus in a center, there are no
openings. The presence of tracheids and/or vessel elements has evolutionary
significance. Vessels (made of vessel members) are more effective; consequently,
more “primitive” plants have more tracheids whereas more “advanced” have more
vessel members. As an example, gymnosperms have only tracheids while most
flowering plants have tracheids and vessel members. Individual development also
mimics this evolutionary trend. Younger flowering plants have more tracheids
whereas mature plants have more vessel members. Primary xylem mostly has
tracheids and vessels with scalariform perforations whereas secondary xylem
(which originates from cambium) consists mostly of vessels with open
perforations. The common name for secondary xylem is wood.

It is a mistake to think that tracheids are better than vessels. In fact, the main
problem is frequently not too slow but too fast water transport. Tracheids have
an advanced connection system (called torus) which has the ability to close pore if
the water pressure is too high and therefore more controllable. Leaking would be
less dangerous in tracheids. And in water-poor environments (like taiga in winter),
plants with tracheids will have the advantage. Contrary, having vessels is like to

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have race car for ordinary life; only flowering plants “learned” how to use them
effectively.

Dead cells are useful but hard to control. However, if xylem transport needs to
be decreased, there is a way. Xylem parenchyma cells will make tyloses
(“stoppers”) which will grow into dead tracheary elements and stop water if
needed. Many broadleaved trees use tyloses to lower xylem transport before the
winter.

The phloem generally occurs adjacent, or right next to, the xylem, with the xylem
facing the inner part of the plant and the phloem facing the outer part of the
plant. The main functions of the phloem are the transportation of sugars and
mechanical support. The four types of phloem cells are: sieve tube cells,
companion cells, fibers (the only dead cells in phloem), and parenchyma. Sieve
tube cells of flowering plants have cytoplasm flowing through perforations (sieve
plates) between cells but do not contain nuclei. Companion cells will make proteins
for them. However, in gymnosperms and more “primitive” plants there are no
companion cells at all, so sieve tube cells do contain nuclei. This is comparable to
red blood cells in vertebrates: while mammals have them anucleate, erythrocytes
of other vertebrates contain nucleus. The secondary phloem generally has more
fibers than the primary phloem.

This small table summarizes differences between xylem and phloem.

Xylem Phloem
Contains mostly Dead cells Living cells
Transport Water Sugar
Direction Up Down
Biomass Big Small

5.1.5 Periderm

Periderm is a secondary dermal tissue which arises inside the stem ground tissue,
closer to the surface. Like the other dermal tissue (epidermis), it is a complex
tissue. It includes three layers (starting from surface): phellem (cork), phellogen
(cork cambium) and phelloderm (Fig. 5.7). Phellem consists of large dead cells with
secondary walls saturated with suberin, and is the main, thickest component of
periderm. Phellogen is a lateral meristem, like cambium; it often arises
fragmentarily (and also temporarily) and does not cover the whole stem
under-surface. But when phellem starts to grow, all peripheral tissues (like
epidermis) will be separated from water transport and eventually die. Phellogen
makes phellem towards the surface, and phelloderm towards the next layer
(phloem). Phelloderm is a minute tissue, and does not play significant role in the
periderm.

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 Figure 5.7. Principal location of stem tissues (simplified).

In older plants, phellogen arises deeper, sometimes inside phloem and separates
outer layers of phloem from vascular cylinder. All this mixture of tissues
(phellogen, phellem, phelloderm, epidermis and upper layers of phloem) considered
as a bark.

5.1.6 Absorption Tissues

Poikilohydric plants do not save water and they can survive even complete
desiccation because their cells will hibernate. An example of a poikilohydric
plants would be mosses. Homoiohydric plants (which are majority of plants2),
however, do save water. They try to support the water content and do not survive
complete desiccation. An example of a homoiohydric plant would be any “typical”
plant, saying, corn. Somehow similar traits are comparable in poikilothermic
animals, such as reptiles, and homoiothermic animals, such as birds and mammals,
except in reference to body heat rather than water conservation.

Absorption tissues are always simple, primary tissues. Most important of them is
rhizodermis (rhizoderm), or root hairs, which originates from protoderm
(proto-epidermis), but its lifespan is much shorter than of epidermis. There are
other absorption tissues, for example, velamen, which originates from the root
cortex and consists of large, empty, easy to get wet dead cells.

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5.1.7 Other Tissues

Secretory tissues spread across the plant body, concentrating in leaves and young
stems. These tissues may secrete latex, volatile oils, mucus and other chemicals.
Its functions can be attraction or dis-attraction, communication or defense, and
many others.

In addition to tissues, plant body may contain idioblasts, cells which are quite
dissimilar from surrounding cells. Idioblasts used for accumulation of unusual
(and possibly dangerous) compounds like myrosinase, protein splitting
glucosinolates into sugars and toxic isothiocyanate (mustard oil). We use mustard
oil as a spice but for the plant, it works like a binary chemical weapon against
insect herbivores: when myrosinase-containing idioblasts are damaged, mustard
oil kills damaging insects. Among plants, the whole order Brassicales from rosids
is capable to produce myrosinase, examples are different cabbages (Brassica
spp.), papaya (Carica), horseradish tree (Moringa) and many others.

5.2 Organs and Organ Systems

Plantae (Vegetabilia) (Fig. 6.1) have three different types of body construction
(Fig. 5.8). The most primitive plants have thallus body, more advanced is the shoot
(unipolar) plant body, and most land plants have the bipolar plant body. The thallus
plant body is flat, similar to leaf but do not differentiated into particular organs.
Most gametophytes (except true mosses) have this type, and also few
sporophytes (which mostly are reduced water plants). Shoot (unipolar) plant body
consists only of branching shoots, roots are absent. This is typical to all
Bryophyta sporophytes, mosses (Bryopsida) gametophytes, and also to
sporophytes of Psilotopsida (whisk ferns). Finally, bipolar plant body has both
shoots and roots (Fig 5.11). Most bipolar plants have shoots consist of stems and
leaves, but this is not an absolute requirement since young plant stems are
normally green and can do photosynthesis.

Figure 5.8. Evolution of plants2 body types: a–e thallus gametophytes, a thallus sporophyte, b–d
shoot sporophyte, e bipolar sporophyte.

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 Figure 5.9. Young seedling with epicotyl and hypocotyl.

Typical organs of bipolar plant are stems (axial aerial organs with continuous
growth), leaves (flat lateral organ with restricted growth), roots (axial soil organ
modified for absorption) and floral units (FU) which are elements of the
generative system (fructifications) such as a pine cone or any flower.

Buds, fruits, seeds and specific to seedlings hypocotyl and epicotyl are
non-organs for different reasons: buds are just young shoots, fruit is the ripe
flower, hypocotyl
is a part of stem between first leaves of the seedling (cotyledons) and root (i.e.,
stem/root transition place), epicotyl is first internode of stem (Fig. 5.9), and
finally, seed is a chimeric structure with three genotypes so it is impossible to
call it “organ”.

Root, stem, leaf and FU are four basic plant organs (Fig. 5.10) which in bipolar
plant could be grouped in root and shoot system; the latter is frequently split into
generative shoot system (bearing FU), and vegetative shoot system (without FU).

Figure 5.10. Bipolar plant:

organ systems and four
organs.

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Vegetative shoot system usually consists of main and secondary shoots; shoots
contain terminal buds, axillary (lateral) buds, stem (nodes and internodes) and
leaves. We will start from leaves.

5.3 The Leaf

The first and ultimate goal of every plant is photosythesis. If a plant is

multicellular, it usually develops relatively large, flat structures which goal is to
catch sun rays. Terrestrial plants are no exception; most probably, they started
to build their body with organs similar to present day leaves.

Figure 5.11. Systems of organs and organs of bipolar plant.

A leaf is lateral photosynthetic organ of shoot with restricted growth. Its

functions are photosynthesis, respiration, transpiration, and synthesis of
secondary chemicals. Features of a leaf (i.e., characters help to distinguish it)

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include having a bud in the axil, not growing by apex, not producing new leaves or
shoots, and having hierarchal morphology.

Figure 5.12. How to distinguish compound leaves (left) from branches (right).

5.3.1 Morphology of the Leaf

Morphology means external, well visible structural features whereas anatomy
needs tools like a microscope and/or scalpel. Leaves are very important in plant
morphology. The ability to describe the leaf is a must even for novices in botany.

In all, plants are fractal organisms, like Sierpinski triange (Fig. 5.13). All fractals
are self-similar (Fig. 5.14), and plants are no exception. Self-similarity, or
“Russian doll effect”means that almost every part of plant may be a part of the
bigger complex, this bigger one—the part of even bigger system, and so on. This is
what we see in leaves as levels of hierarchy. Simple leaves have just one level of
hierarchy whereas compound leaves have two or more levels of hierarchy.
Compound leaves are sometimes mixed with branches but there are many other
characteristics which allow to distinguish them (Fig. 5.12)

Figure 5.13. One of simple fractals: Sierpinski triangle.

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 Figure 5.14. The example of self-similarity.

To describe leaves, one should always note the level of hierarchy like “on the first
level of hierarchy, the shape is..., on the second level of hierarchy, the shape is ...”
As it was mentioned above, leaf hierarchy is similar to Russian dolls: every smaller
doll has a bigger doll (next hierarchy level) outside. For example, if the leaf is
compound (consists of multiple leaflets), the overall shape of it could be, saying,
round (circular) but the shape of individual leaflet of the very same leaf could be
ovate (Fig. 5.15). As a result, the description will say that on first level of
hierarchy the leaf is ovate, and on the third level — circular.

There are three types of leaf characters: general, terminal, and repetitive.
General characters are only applicable to the whole leaf. Terminal characters are
only applicable to the terminal leaflets. Terminals are the end parts of leaves,
they do not split in smaller terminals; clover leaf, for example, has 3 terminals.
Lastly, repetitive characters repeat on each level of leaf hierarchy. General and
terminal characters do not depend on hierarchy. Repetitive characters may be
different on each step of hierarchy.

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Figure 5.15. Leaves with one, two and three levels of hierarchy. Please note that the last leaf is
ovate on the first and second level but circular on the third level of hierarchy.

General characters of leaf include stipules and other structures located near leaf
base (Fig. 5.16): sheath (typical for grasses and other liliids) and ocrea (typical
forbuckwheat family, Polygonaceae).

Figure 5.16. From left to right: sheath, stipules and ocrea.

Repetitive characters are the shape of the leaf (Fig.5.17), leaf dissection, and
whether the blade is stalked (has petiole) or not.

Terminal characters are applicable only to terminal leaflets of leaves. These

characters (Fig. 5.19) are the shape of the leaf blade base, the leaf tip, the type
of margin, the surface, and the venation. The base of the leaf blade could be
rounded, truncate (straight), cuneate, and cordate. The leaf apex could be
rounded, mucronate, acute, obtuse, and acuminate. Leaf margin variants are
entire (smooth) and toothed: dentate, serrate, double serrate and crenate.

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Figure 5.17. Leaf shapes.

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Figure 5.18. Leaf dissection.

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 Figure 5.19. Terminal leaf characters.

Leaf veins are vascular bundles coming to the leaf from stem. Frequently, there is
a main vein and lateral veins (veins of second order). There are multiple
classifications of leaf venation; and example is shown on the Figure 5.20.

Note that in dichotomous venation, each vein divides into two similar parts which
is known as dichotomous branching. The example of dichotomous venation is the
leaf of maidenhair tree, ginkgo (Ginkgo biloba). Another frequently segregated
type of venation is parallellodromous, but in essence, this is acrodromous
venation in linear leaves (for example, leaves of grasses) where most of veins are
almost parallel.

To characterize the whole leaf, one might use the following plan:

1. General characters (leaf as a whole):

(a) stipules (present / absent, deciduous / not, how many, size, shape);

(b) base (sheath / no sheath, ocrea / no ocrea)

2. First level of hierarchy: repetitive characters:

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 (a) symmetry (symmetrical / asymmetrical);

(b) shape;

(c) dissection;

(d) petiole (presence and length)

3. Second level of hierarchy

4. Third level of hierarchy, and so on

5. Terminal characters (leaflets):

(a) base of leaf blade (rounded, truncate, cuneate, cordate)

(b) apex (rounded, mucronate, acute, obtuse, acuminate);

(c) margin (whole, dentate, serrate, double serrate, crenate );

(d) surfaces (color, hairs etc.);

(e) venation (apo-, hypho-, acro-, ptero-, actinodromous)

Figure 5.20. The simple classification of leaf venation.

Heterophylly refers to a plant having more than one kind of leaf. A plant can have
both juvenile leaves and adult leaves, water leaves and air leaves, or sun leaves
and shade leaves. A leaf mosaics refers to the distribution of leaves in a single
plane perpendicular to light rays, this provides the least amount of shading for
each leaf.

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Leaves have seasonal lives; they arise from the SAM through leaf primordia, and
grow via marginal meristems. The old leaves separate from the plant with an
abscission zone.

The famous poet and writer Johann Wolfgang Goethe is also considered a founder
of plant morphology. He is invented an idea of a “primordial plant” which he called
“Urpflanze” where all organs were modifications of several primordial ones. In
accordance to Goethe’s ideas, plant morphology considers that many visible plant
parts are just modifications of basic plant organs.

Modifications of the leaf include spines or scales for defense, tendrils for
support, traps, “sticky tapes”, or urns for interactions (in that case, catching
insects), plantlets for expansion, and succulent leaves for storage. Plantlets are
little mini plants that grow on the main plant and then fall off and grow into new
plants; the most known example is Kalanchoe (“mother of thousands”) which
frequently uses plantlets to reproduce. Plants that have insect traps of various
kinds are called carnivorous plants (in fact, they are still photoautotrophs and use
insect bodied only as fertilizer). Several types of these are the cobra lily
(Darlingtonia), various pitcher plants (Nepenthes, Cephalotus, Sarracenia), the
butterwort (Utricularia), the sundew (Drosera), and the best known, the Venus
flytrap (Dionaea).

5.3.2 Anatomy of the Leaf

Anatomically, leaves consist of epidermis with stomata, mesophyll (kind of

parenchyma) and vascular bundles, or veins (Fig. 5.22). The mesophyll, in turn, has
palisade and spongy variants. Palisade mesophyll is located in the upper layer and
serves to decrease the intensity of sunlight for the spongy mesophyll, and also
catches slanted sun rays. The palisade mesophyll consists of long, thin, tightly
arranged cells with chloroplasts mostly along the sides. The spongy mesophyll
cells are roughly packed, they are rounded and have multiple chloroplasts (Fig.
5.21).

Figure 5.21.
Leaf anatomy.

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When a typical stem vascular bundle (which has xylem under phloem) enters the
leaf, xylem usually faces upwards, whereas phloem faces downwards. Bundles of
C4- plants have additional bundle sheath cells in their vascular bundles.

The epidermis includes typical epidermal cells, stomata surrounded with guard
cells (also optionally with subsidiary cells), and trichomes. Almost all epidermal
cells are covered with waterproof cuticle, rich of lignin and waxes.

The stomata assists in gas exchange, cooling and water transpiration. There are
two guard cells paired together on each side of the stoma. These guard cells are
kidney beans shaped and have a thicker cell wall in the middle. The thicker cell
wall on the inside makes use of the so-called “bacon effect” (when bacon slice
curved on the frying pan) because thinner part of the cell wall is more flexible
and therefore bends easier. The same curving effect might be seen in blowing air
balloon with the piece of scotch on one side. The opening of the stoma starts
from K+ accumulation, then osmosis inflates guard cells, and finally the uneven cell
wall facilitates the opening of stoma. The stoma closes when the potassium ions
exit the cell and water amount decreases in its vacuoles (Fig 5.23).

Figure 5.22. Left to right, top to bottom: leaf of sclerophyte Pinus, leaf of salt-avoiding
(succulent-like) halophyte Salsola (epidermis is at the bottom), shade leaf of Sambucus, leaf of
Syringa with guards cells (bottom left). Magnifications ×100 (first) and ×400 (others).

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 Figure 5.23. Closed and opened stoma. Cell walls are white, cytoplasm green, vacuoles blue.

In most cases, the lower epidermis contains more stomata than the upper
epidermis because the bottom of the leaf is cooler and transpiration there is
safer. A similar logic is applicable to trichomes (hairs): they are also more
frequent on the lower side of the leaf.

5.3.3 Ecological Forms of Plants

When plants adapt to the particular environment conditions, leaves usually

respond first. Conversely, one can estimate the ecology of plant simply looking on
its leaves.

In regards to water, there are four main types of plants: xerophytes, mesophytes,
hygrophytes, and hydrophytes. Xerophytes are adapted to the scarce water (Fig.
5.22), they could be sclerophytes (usually with prickly and/or rich of
sclerenchyma leaves) and succulents (with water-accumulating stems or leaves).
Mesophytes are typical plants which adapt to regular water. Hygrophytes live in
constantly wet environment, their leaves adapted to high transpiration and
sometimes even to guttation (excretion of water drops). Hydrophytes grow in
water, their leaves are frequently highly dissected to access more gases
dissolved in water, and their leaf petioles and stems have air canals to supply
underwater organs with gases.

In regards to light, plants could be sciophytes or heliophytes. Sciophytes prefer

the shade to sunlight, their leaves contain mostly spongy mesophyll. Heliophytes
prefer the full sun and therefore have leaves filled with palisade mesophyll. The
intermediate group are “partial shade” plants.

Halophytes, nitrate halophytes, oxylophytes, and calciphytes are ecological

groups adapted to the over-presence of particular chemicals. Halophyte plants
are frequent, they accumulate (and look similarly to succulents), excrete or avoid
(which looks like sclerophyte) sodium chloride (NaCl). They grow in salty places:
sea shores, salt deserts and solonets prairies. Nitrate halophyte plants grow on

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soils rich in NaNO3. Oxylophytes grow in acidic soils, whereas calciphytes grow in
basic, chalk soils rich in CaCO3.

Leaves will also reflect adaptations to the substrate, ecological forms named
psammophytes (grow on sand), petrophytes (grow on rocks), and rheophytes
(grow in fast springs). The latter plants frequently have serious simplifications in
their body plan, their leaves and stems are often reduced to form a thallus-like
body.

Parasitic plants could be classified in mycoparasites, hemiparasites, and

phytoparasites. Mycoparasitic plants feed on soil fungi, phytoparasitic plants are
either plant root parasites or plant stem parasites lacking chlorophyll and
photosynthesis. Hemiparasitic plants are those which still have chloroplasts but
take the significant part of water and even organic compounds from the host
plant (like mistletoe, Viscum).

5.4 The Stem

The stem is an axial organ of shoot. It has functions of support, transportation,

photosynthesis, and storage. Stem has radial structure, no root hairs and grows
continuously.

5.4.1 Morphology of the Stem

Stem morphology is simple. Its components are nodes (places where leaves
are/were attached) and internodes, long or short (in the last case, plant
sometimes appears to be stemless, rosette-like).

Stems are different by the type of phyllotaxy. The phyllotaxy refers to the
arrangement of leaves. If there is one leaf per node, it is a spiral (alternate)
arrangement. Two leaves per node means opposite arrangement. Opposite leaves
can be all in the same plane or each pair can rotate at 900. If there are more
than two leaves per node, it is a whorled arrangement, and each whorl can also
rotate.

Each type of spiral phyllotaxy has its own angle of divergence. Multiple types of
spiral leaf arrangement mostly follow the Fibonacci sequence:

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Figure 5.24. Types of phyllotaxy (leaf arrangement): a spiral (alternate), b and c opposite, d
whorled.

This sequence of numbers made with simple rule: in the every following fraction,
the numerator and denominator are sums of two previous numerators and
denominators, respectively. The sequence looks fairly theoretical but amazingly,
it is fully applicable to plant science, namely to different types of spiral
phyllotaxy (Fig. 5.25).

Figure 5.25. Four first Fibonacci types of spiral phyllotaxy: 1/2 , 1/3 , 2/5, and 3/8 .

To determine formula of spiral phyllotaxy, one needs to start with arbitrary leaf
(or leaf scar) and then find the next (upper) one which is directed the same way,
lays on the same virtual line. Then, the imaginary spiral should be drawn trough
basements from the started leaf to the corresponding upper leaf.

This spiral should go through all intermediary leaves, there might be one, two or
more intermediary leaves. Also, the spiral will go at least one time around the
stem. (Instead of the imaginary spiral, it is sensible to use a thin thread). One
needs to count all leaves in the spiral except the first, and also count number of
rotations.

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The number of leaves counted will be the denominator of the formula, and the
number of rotations is the numerator. This is how Fibonacci numbers appear in
plant morphology.

These phyllotaxy formulas are relatively stable and sometimes even

taxon-specific. For example, grasses (Gramineae) have 1/2 phyllotaxy, sedges
(Carex) 1/3, many Rosaceae (like apple, Malus or cherry, Prunus) have 2/5, willows
frequently have 3/8, etcetera.

It is still not absolutely clear why the spiral phyllotaxy is under such a theoretical
mathematical rule. The most feasible hypothesis emphasizes mathematical
problem of circle packing and the competition between leaf primordia around
SAM.

5.4.2 Anatomy of the Primary Stem

Plant evolution resulted first in the primary stems with no lateral meristems and
secondary tissues. Only long after plants “learned” how to thicken their stems.

Figure 5.26. Developmental origin of stem tissues (simplified). Letters e, p, a show respectively
where endoderm, pericycle and vascular cambium might appear.

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Development of stem starts from stem apical meristem (SAM) on the top of plant.
The SAM produces three primary meristems: procambium, protoderm, and
ground meristem. Protoderm cells differentiate into epidermal cells. The ground
meristem differentiates into the cortex and pith. The procambium raises
between the cortex and the pith. It forms vascular bundles or vascular cylinder
(Fig. 5.26).

Figure 5.27. Developmental origin of stem tissues (detailed). Root tissues have similar ways of
development

The outer layers of the procambium form the primary phloem. The inner layers
become the primary xylem. The middle layer can be entirely spent or will make
cambium for the secondary thickening. At times, the layers of the outside of the
procambium can form a pericycle. Sometimes the innermost layer of the cortex
can form an endodermis (endoderm) (Fig. 5.28), and outermost layer makes the
exodermis (exoderm). All these layers are some kind of the “border control”
between functionally different layers of stem. Another frequent variant is the
development of collenchyma in the cortex adjacent to epidermis.

Vascular bundles connect leaves and stems. In many plants, they form a ring on
the cross-section of the stem. Parenchyma (ground tissue) between vascular
bundles typically belongs to both cortex and pith. Another variant is a vascular
cylinder, structure which fully encircles the stem. Liliid (monocot) stems
generally have dispersed vascular bundles. These three variants are steles,
overall configurations of the primary vascular system of the plant stem (Fig.
5.29). The most frequent kinds of steles are eustele (vascular bundles in a ring),
solenostele (vascular cylinder) and ataktostele (dispersed vascular bundles).

All these types were probably originated from protostele, configuration where
central xylem is surrounded with phloem and no pith is present (Fig. 5.30). While
the protostele was typical for many prehistoric plants, now only some lycophytes

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(Huperzia) have protostele in stems. Saying that, it is important to note that
roots of most plants have vascular tissues arranged similarly to protostele.

Figure 5.28. Anatomy of the primary stem (right). Slanted font is used for “optional” tissues. Small
image on the left is the young stem consisted of epidermis, cortex, procambium and pith.

5.5 The Root

Root is a latest evolutionary innovation in the vegetative plant anatomy. Many

“primitive” plants (all mosses and even some ferns like Psilotum) do not have roots;
some flowering water plants like the¿ rootless duckweed (Wolffia) or the coontail
(Ceratophyllum) have also reduced their roots. However, large homoiohydric
plants need the constant supply of water and minerals, and this evolutionary
challenge was responded with appearance of the root system.

Root in an axial organ of plant with geotropic growth. One of root functions is to
supply anchorage of the plant body in soil or on various surfaces. Other functions
include water and mineral absorption and transport, food storage, and
communication with other plants.

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Figure 5.29. Left to right, top to bottom: eustele (stem segment), xylem vessel (longitudinal
section) and ataktostele (stem segment and vascular bundle). First photo is from the stem of
Helianthus, second from Trifolium, last two from the stem of Zea. Magnifications ×100 (first and
third) and ×400 (second and fourth)

Figure 5.30. Steles (left to right): protostele, solenostele, eustele, ataktostele. Xylem is lined,
phloem is dotted.

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5.5.1 Morphology of the Root

There are two types of root systems. The first is a fibrous root system which has
multiple big roots that branch and form a dense mass which does not have a
visible primary root (“grass-like”). The other is the tap root system which has one
main root that has branching into lateral roots (“carrot-like”).

Along with having different systems, there are different types of roots: primary
root originated from the root of the seedling, secondary (lateral) roots originate
from the primary roots, and adventitious roots originate on stems (sometimes
also on leaves), the example are prop roots of screw pine (Pandanus).

Roots employ many different modifications which help to protect, interact and
storage. For example, roots of parasitic plants are modified into haustoria which
sink themselves into the vascular tissue of a host plant and live off of the host
plant’s water and nutrients.

Roots of mangroves (plants growing in ocean coastal swamps) are frequently

modified into supportive aerial roots (“legs”). Since these swamp plants need
oxygen to allow cell respiration in underground parts, there are pneumatophores,
specialized roots which grow upward (!) and passively catch the air via multiple
pores. Plants which grow on sand (psammophytes, see above) have another
problem: their substrate constantly disappears. To avoid this, plants developed
contractile roots which may shorten and pull plant body deeper into the sand.

Some orchid roots are green and photosynthetic (Fig. 5.31)! However, as a rule,
root is the heterotrophic organ, because root cells have no access to the light.

Root nodules present on the roots of nitrogen-fixing plants, they contain bacteria
capable to deoxidize athmospheric nitrogen into ammonia: N2 → NH3. Root
nodules contain also hemoglobin-like proteins which facilitate nitrogen fixation by
keeping oxygen concentration low. Nitrogen-fixing plants are especially frequent
among faboid rosids: legumes (Leguminosae family) and many other genera (like
alder, Alnus, or Shepherdia, buffaloberry) have root nodules with bacteria. Some
other plants (mosquito fern, Azolla and dinosaur plant, Gunnera) employ
cyanobacteria for the same purpose.

Figure 5.31. Photosynthetic roots of leafless

orchid Chiloschista segawai.

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Mycorrhiza is a root modification started when fungus penetrates root and
makes it more efficient in mineral and water absorption: it will exchange these
for organic compounds. In addition to mycorrhizal fungi, endophytic fungi inhabit
other plant organs and tissues.

5.5.2 Anatomy of the Root

On the longitudinal section of young growing root, there are different horizontal
layers, zones: root cap covering division zone, elongation zone, absorption zone,
and maturation zone (Fig. 5.32). The root cap protects the root apical meristem
(RAM), which is a group of small regularly shaped cells. A small, centrally located
part of the RAM is the quiescent center where initial cells divide and produce all
other cells of root Root cap is responsible for the geotropic growth, if the root
tip comes into contact with a barrier, root cap will feel it and will grow on a
different direction to go around it.

The elongation zone is where the cells start to elongate, giving it length. The
absorption zone is where the rhizodermis tissue (root hairs) develops and where
water and nutrients are absorbed and brought into the plant. Within the
maturation zone, root hairs degrade, many cells start to acquire secondary walls
and lateral roots develop (Fig. 5.32).

Figure 5.32. Root zones: 0 root cap, 1 division zone, 2 elongation zone, 3 absorption zone, 4
maturation zone.

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On the cross-section of the root made within absorption zone, the first tissue is
the rhizodermis, which is also known as the root epidermis, then cortex, which
segregates external exodermis and internal endodermis one-cellular layers, and
vascular cylinder (Fig. 5.33). Typically, roots have no pith. In some cases (for
example, in orchids), cortex may give multi-layered velamen (see above), another
absorption tissue.

Figure 5.33. Left to right, top to bottom: Ranunculus root with 4-rayed xylem, Salix root with the
lateral root developing, Smilax root with visible Casparian stripes in the endodermis; Zea root
longitudinal section with root cap, division and elongation zones. Magnifications ×100 (second) and
×40 (others).

Vascular cylinder is located in the center of the root, it contains the pericycle
which is made of mostly parenchyma and bordering endodermis. Pericycle cells
may be used for storage, they contribute to the vascular cambium, and initiate
the development of lateral roots. Consequently, lateral roots are developing
endogenously and break tissues located outside, like aliens in the famous movie.
Root phloem is arranged in several strands whereas xylem typically has a radial,

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sometimes star-shaped structure with few rays (Fig. 5.34). In the last case,
phloem strands are located between rays of xylem.

Root tissues develop in the way similar to stem, RAM gave rise to ground
meristem, procambium, and the protoderm, which in turn make all primary tissues
mentioned above. Later, pericycle develops into lateral roots or the vascular
cambium which in turn produces into the secondary xylem and phloem. The
secondary root is similar to secondary stem.

Figure 5.34. Anatomy of root: cross-section through the maturation zone.

5.5.3 Water and Sugar Transportation in Plants

Plants need water to supply photosynthesis (the oxygen is from water!), to cool
down via transpiration, and to utilize diluted microelements. Dead velamen
(paper-like), rhizoids (hair-like), and living rhizodermis (rhizoderm) are
responsible for water uptake.

In rhizodermis, root hairs increase the surface area where the plant has to
absorb the nutrients and water. To take water, hair cells increase concentration
of organic chemicals (the process which needs ATP) and then use osmosis. There
are two ways that water transport may go: apoplastic or symplastic. Apoplastic
transport moves water through the cell walls of cortex: from the rhizodermis to
the endodermis. Endodermis cell walls bear Casparian strips (rich of hydrophobic
suberin and lignin) which prevent the water from passing through the cell wall and
force symplastic transport (Fig 5.35) through cytoplasms and plasmodesmata.
Symplastic transport there is directed to the center of root only and requires
ATP to be spend.

By pumping water inside vascular cylinder and not letting it back, endodermis cells
create the root pressure (Fig. 5.36). It is easy to observe on tall herbaceous
plants cut near the ground: drops of water will immediately appear on the cutting.
Inside tracheary elements of xylem, water moves with the root pressure,
capillary force and the sucking pressure of transpiration. The latter means that
water column does not want to break and if water disappears from the top

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(stomata on leaves), it will move water inside plant. The main direction of water
movement is from roots to leaves, i.e. upwards.

Figure 5.35. Symplastic and apoplastic transport in root.

Figure 5.36. The origin of root pressure: water comes into vascular cylinder but cannot go ack
because of endoderm (brown line). The only possible way is to go up.

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Products of photosynthesis (sugars) are moving inside living cells of phloem; these
cells (sieve tubes) use only symplastic transport to distribute glucose and other
organic compounds among all organs of plants. In fact, phloem transports these
components in all directions: to the flowers (usually upwards), and at the same
time to the roots (usually downwards).

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